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Breeding for early or late maturation is certainly a reality; it is also possible to breed for fast or slow flowering and even or sequential ripening. In general, crosses between early-maturing plants give rise to early-maturing offspring, crosses between late-maturing plants give rise to late maturing offspring, and crosses between late- and early maturing plants give rise to offspring of intermediate maturation. This seems to indicate that maturation of Cannabis is not controlled by the simple dominance and recessiveness of one gene but probably results from incomplete dominance and a combination of genes for separate aspects of maturation. For instance, Sorghum maturation is controlled by four separate genes. The sum of these genes produces a certain phenotype for maturation. Although breeders do not know the action of each specific gene, they still can breed for the total of these traits and achieve results more nearly approaching the goal of timely maturation than the parental strains.
g) Root Production The size and shape of Cannabis root systems vary greatly. Although every embryo sends out a taproot from which lateral roots grow, the individual growth pattern and final size and shape of the roots vary considerably. Some plants send out a deep taproot, up to 1 meter (39 inches) long, which helps support the plant against winds and rain. Most Cannabis plants, however, produce a poor taproot which rarely extends more than 30 centimeters (1 foot). Lateral growth is responsible for most of the roots in Cannabis plants. These fine lateral roots offer the plant additional support but their primary function is to absorb water and nutrients from the soil. A large root system will be able to feed and support a large plant. Most lateral roots grow near the surface of the soil where there is more water, more oxygen, and more available nutrients. Breeding for root size and shape may prove beneficial for the production of large rain- and wind resistant strains. Often Cannabis plants, even very large ones, have very small and sensitive root systems. Recently, certain alkaloids have been discovered in the roots of Cannabis that might have some medical value. If this proves the case, Cannabis may be cultivated and bred for high alkaloid levels in the roots to be used in the commercial production of pharmaceuticals. As with many traits, it is difficult to make selections for root types until the parents are harvested. Because of this many crosses are made early and seeds selected later.
h) Branching The branching pattern of a Cannabis plant is determined by the frequency of nodes along each branch and the extent of branching at each node. For examples, consider a tall, thin plant with slender limbs made up of long internodes and nodes with little branching (Oaxaca, Mexico strain). Compare this with a stout, densely branched plant with limbs of short internodes and highly branched nodes (Hindu Kush hashish strains). Different branching patterns are preferred for the different agricultural applications of fiber, flower, or resin production. Tall, thin plants with long internodes and no branching are best adapted to fiber production; a short, broad plant with short internodes and well developed branching is best adapted to floral production. Branching structure is selected that will tolerate heavy rains and high winds without breaking. This is quite advantageous to outdoor growers in temperate zones with short seasons. Some breeders select tall, limber plants (Mexico) which bend in the wind; others select short, stiff plants (Hindu Kush) which resist the weight of water without bending.
i) Sex Attempts to breed offspring of only one sexual type have led to more misunderstanding than any otherfacet of Cannabis genetics. The discoveries of McPhee (1925) and Schaffner (192 showed that pure sexual type and hermaphrodite conditions are inherited and that the percentage of sexual types could be altered by crossing with certain hermaphrodites. Since then it has generally been assumed by researchers and breeders that a cross between ANY unselected hermaphrodite plant and a pistillate seed-parent should result in a population of all pistillate offspring. This is not the case. In most cases, the offspring of hermaphrodite parents tend toward hermaphrodism, which is largely unfavorable for the production of Cannabis other than fiber hemp. This is not to say that there is no tendency for hermaphrodite crosses to alter sex ratios in the offspring. The accidental release of some pollen fro predominantly pistillate hermaphrodites, along with the complete eradication of nearly every staminate and staminate hermaphrodite plant may have led to a shift in sexual ratio in domestic populations of sinsemilla Cannabis. It is commonly observed that these strains tend toward 60% to 80% pistillate plants and a few pistillate hermaphrodites are not uncommon in these populations.
However, a cross can be made which will produce nearly all pistillate or staminate individuals. If the proper pistillate hermaphrodite plant is selected as the pollen parent and a pure pistillate plant is selected as the seed parent it is possible to produce an F1, and subsequent generations, of nearly all pistillate offspring. The proper pistillate hermaphrodite pollen-parent is one which has grown as a pure pistillate plant and at the end of the season, or under artificial environmental stress, begins to develop a very few staminate flowers. If pollen from these few staminate flowers forming on a pistillate plant is applied to a pure pistillate seed parent, the resulting F1 generation should be almost all pistillate with only a few pistillate hermaphrodites. This will also be the case if the selected pistillate hermaphrodite pollen source is selfed and bears its own seeds. Remember that a selfed hermaphrodite gives rise to more hermaphrodites, but a selfed pistillate plant that has given rise to a limited number of staminate flowers in response to environmental stresses should give rise to nearly all pistillate offspring. The F1 offspring may have a slight tendency to produce a few staminate flowers under further environmental stress and these are used to produce F2 seed. A monoecious strain produces 95+% plants with many pistillate and staminate flowers, but a dioecious strain produces 95+% pure pistillate or staminate plants. A plant from a dioecious strain with a few intersexual flowers is a pistillate or staminate hermaphrodite. Therefore, the difference between monoecism and hermaphrodism is one of degree, determined by genetics and environment.
Crosses may also be performed to produce nearly all staminate offspring. This is accomplished by crossing a pure staminate plant with a staminate plant that has produced a few pistillate flowers due to environmental stress, or selfing the latter plant. It is readily apparent that in the wild this is not a likely possibility. Very few staminate plants live long enough to produce pistillate flowers, and when this does happen the number of seeds produced is limited to the few pistillate flowers that occur. In the case of a pistillate hermaphrodite, it may produce only a few staminate flowers, but each of these may produce thou sands of pollen grains, any one of which may fertilize one of the plentiful pistillate flowers, producing a seed. This is another reason that natural Cannabis populations tend toward predominantly pistillate and pistillate hermaphrodite plants. Artificial hermaphrodites can be produced by hormone sprays, mutilation, and altered light cycles. These should prove most useful for fixing traits and sexual type. Drug strains are selected for strong dioecious tendencies. Some breeders select strains with a sex ratio more nearly approaching one than a strain with a high pistillate sex ratio. They believe this reduces the chances of pistillate plants turning hermaphrodite later in the season.
2. Seedling Traits
Seedling traits can be very useful in the efficient and purposeful selection of future parental stock. If accurateselection can be exercised on small seedlings, much larger populations can be grown for initial selection, as less space is required to raise small seedlings than mature plants. Whorled phyllotaxy and resistance to damping-off are two traits that may be selected just after emergence of the embryo from the soil. Early selection for vigor, hardiness, resistance, and general growth form may be made when the seedlings are from 30 to 90 centimeters (1 to 3 feet) tall. Leaf type, height, and branching are other criteria for early selection. These early-selected plants cannot be bred until they mature, but selection is the primary and most important step in plant improvement. Whorled phyllotaxy is associated with subsequent anomalies in the growth cycle (i.e., multiple leaflets and flattened or clubbed stems). Also, most whorled plants are staminate and whorled phyllotaxy may be sex-linked.
3. Leaf Traits
Leaf traits vary greatly from strain to strain. In addition to these regularly occurring variations in leaves, there are a number of mutations and possible traits in leaf shape. It may turn out that leaf shape is correlated with other traits in Cannabis. Broad leaflets might be associated with a low calyx-to-leaf ratio and narrow leaflets might be associated with a high calyx-to-leaf ratio. If this is the case, early selection of seedlings by leaflet shape could determine the character of the flowering clusters at harvest. Both compound and webbed leaf variations seem to be hereditary, as are general leaf characteristics. A breeder may wish to develop a unique leaf shape for an ornamental strain or increase leaf yield for pulp production.
A peculiar leaf mutation was reported from an F1 Colombian plant in which two leaves on the plant, at the time of flowering, developed floral clusters of 5-10 pistillate calyxes at the intersection of the leaflet array and the petiole attachment, on the adaxial (top) side of the leaf. One of these clusters developed a partial staminate flower but fertilization was unsuccessful. It is unknown if this mutation is hereditary. From Afghanistan, another example has been observed with several small floral clusters along the petioles of many of the large primary leaves.
4. Fiber Traits
More advanced breeding has occurred in fiber strains than any other type of Cannabis. Over the years many strains have been developed with improved maturation, increased fiber content, and improved fiber quality as regards length, strength, and suppleness. Extensive breeding programs have been carried on in France, Italy, Russia, and the United States to develop better varieties of fiber Cannabis. Tall limbless strains that are monoecious are most desirable. Monoeciousness is favored, because in dioecious populations the staminate plants will mature first and the fibers will become brittle before the pistillate plants are ready for harvest. The fiber strains of Europe are divided into northern and southern varieties. The latter require higher temperatures and a longer vegetative period and as a result grow taller and yield more fiber.
5. Floral Traits
Many individual traits determine the floral charactistics of Cannabis This section will focus on the individual traits of pistillate floral clusters with occasional comments about similar traits in staminate floral clusters. Pistillate flowering clusters are the seed-producing organs of Cannabis; they remain on the plant and go through many changes that cannot be compared to staminate plants.
a) Shape The basic shape of a floral cluster is determined by the internode lengths along the main floral axis and within individual floral clusters. Dense, long clusters result when internodes are short along a long floral axis and there are short internodes within the individual compact floral clusters (Hindu Kush). Airy clusters result when a plant forms a stretched floral axis with long internodes between well-branched individual floral clusters (Thailand).
The shape of a floral cluster is also determined by the general growth habit of the plant. Among domestic Cannabis phenotypes, for instance, it is obvious that floral clusters from a creeper phenotype plant will curve upwards at the end, and floral clusters from the huge upright phenotype will have long, straight floral clusters of various shapes.Early in the winter, many strains begin to stretch and cease calyx production in preparation for rejuvenation and subsequent vegetative growth in the spring. Staminate plants also exhibit variation in floral clusters. Some plants have tight clusters of staminate calyxes resembling inverted grapes (Hindu Kush) and others have long, hanging groups of flowers on long, exposed, leafless branches (Thailand).
b) Form The form of a floral cluster is determined by the numbers and relative proportions of calyxes and flowers. A leafy floral cluster might be 70% leaves and have a calyx-to-leaf ratio of 1-to-4. It is obvious that strains with a high calyx-to-leaf ratio are more adapted to calyx production, and therefore, to resin production. This factor could be advantageous in characterizing plants as future parents of bud strains. At this point it must be noted that pistillate floral clusters are made up of a number of distinct parts. They include stems, occasional seeds, calyxes, inner leaves subtending calyx pairs (small, resinous, 1-3 leaflets), and outer leaves subtending entire floral clusters (larger, little resin, 3-11 leaflets). The ratios (by dry weight) of these various portions vary by strain, degree of pollination, and maturity of the floral clusters. Maturation is a reaction to environmental change, and the degree of maturity reached is subject to climatic limits as well as breeders preference. Because of this interplay between environment and genetics in the control of floral form it is often difficultto breed Cannabis for floral characteristics. A thorough knowledge of the way a strain matures is important in separating possible inherited traits of floral clusters from acquired traits. Chapter IV, Maturation and Harvesting of Cannabis, delves into the secrets and theories of maturation. For now, we will assume that the following traits are described from fully mature floral clusters (peak floralstage) before any decline.
c) Calyx Size Mature calyxes range in size from 2 to 12 millimeters (1/16 to 3/8 inch) in length. Calyx size is largely dependent upon age and maturity. Calyx size of a floral cluster is best expressed as the average length of the mature viable calyxes. Calyxes are still considered viable if both pistils appear fresh and have not begun to curl or change colors. At this time, the calyx is relatively straight and has not begun to swell with resin and change shape as it will when the pistils die. It is generally agreed that the production of large calyxes is often as important in determining the psychoactivity of a strain as the quantity of calyxes produced. Hindu Kush, Thai, and Mexican strains are some of the most psychoactive strains, and they are often characterized by large calyxes and seeds. Calyx size appears to be an inherited trait in Cannabis. Completely acclimatized hybrid strains usually have many rather small calyxes, while imported strains with large calyxes retain that size when inbred. Initial selection of large seeds increases the chance that offspring will be of the large-calyx variety. Aberrant calyx development occasionally results in double or fused calyxes, both of which may set seed. This phenomenon is most pronounced in strains from Thailand and India.
d) Color The perception and interpretation of color in Cannabis floral clusters is heavily influenced by the imagination of the cultivator or breeder. A gold strain does not appear metallic any more than a red strain resembles a fire engine. Cannabis floral clusters are basically green, but changes may take place later in the season which alter the color to include various shades. The intense green of chlorophyll usually masks the color of accessory pigments, Chlorophyll tends to break down late in the season and anthocyanin pigments also contained in the tissues are unmasked and allowed to show through. Purple, resulting from anthocyanin accumulation, is the most common color in living Cannabis, other than green. This color modification is usually triggered by seasonal change, much as the leaves of many deciduous trees change color in the fall. This does not mean, however, that expression of color is controlled by environment alone and is not an inheritable trait. For purple color to develop upon maturation, a strain must have the genetically controlled metabolic potential to produce anthocyanin pigments coupled with a responsiveness to environmental change such that anthocyanin pigments are unmasked and become visible. This also means that a strain could have the genes for expression of purple color but the color might never be expressed if the environmental conditions did not trigger anthocyanin pigmentation or chlorophyll breakdown. Colombian and Hindu Kush strains often develop purple coloration year after year when subjected to low night temperatures during maturation. Color changes will be discussed in more detail in Chapter IV Maturation and Harvesting of Cannabis.
Carotenoid pigments are largely responsible for the yellow, orange, red, and brown colors of Cannabis. They also begin to show in the leaves and calyxes of certain strains as the masking green chlorophyll color fades upon maturation. Gold strains are those which tend to reveal underlying yellow and orange pigments as they mature. Red strains are usually closer to reddish brown in color, although certain carotenoid and anthocyanin pigments are nearly red and localized streaks of these colors occasionally appear in the petioles of very old floral clusters. Red color in pressed, imported tops is often a result of masses of reddish brown dried pistils.
Several different portions of floral cluster anatomy may change colors, and it is possible that different genes may control the coloring of these various parts.
The petioles, adaxial (top) surfaces, and abaxial (bottom) surfaces of leaves, as well as the stems, calyxes, and pistils color differently in various strains. Since most of the outer leaves are removed during manicuring, the color expressed by the calyxes and inner leaves during the late flowering stages will be all that remains in the final product. This is why strains are only considered to be truly purple or gold if the calyxes maintain those colors when dried. Anthocyanin accumulation in the stems is sometimes considered a sign of phosphorus deficiency but in most situations results from unharmful excesses of phosphorus or it is a genetic trait. Also, cold temperatures might interfere with phosphorus uptake resulting in a deficiency. Pistils in Hindu Kush strains are quite often magenta or pink in color when they first appear. They are viable at this time and turn reddish brown when they wither, as in most strains. Purple coloration usually indicates that pistillate plants are over-mature and cannabinoid biosynthesis is slowing down during cold autumn weather.”
